Sablefish ( Anoplopoma fimbria ) produce high frequency rasp sounds with frequency modulation

Abstract

Sablefish sounds, named rasps, were recorded at two captive facilities in British Columbia and Washington State. Rasps consisted of highly variable broadband trains of 2 to 336 ticks that lasted between 74 and 10 500 ms. The 260 rasps that were measured contained frequencies between 344 and 34 000 Hz with an average peak frequency of 3409 Hz. The frequency structure of ticks within rasps was highly variable and included both positive and negative trends. This finding makes sablefish one of the few deep-sea fish for which sounds have been validated and described. The documentation of sablefish sounds will enable the use of passive acoustic monitoring methods in fisheries and ecological studies of this commercially important deep-sea fish.

Full text

Sablefish (Anoplopoma fimbria) produce high frequency rasp
sounds with frequency modulation
a)
Amalis Riera,
b)
Rodney A. Rountree, Lucas Agagnier, and Francis Juanes
Department of Biology, University of Victoria, 3800 Finnerty Road, Victoria, BC V8P 5C2, Canada
ABSTRACT:
Sablefish sounds, named rasps, were recorded at two captive facilities in British Columbia and Washington State.
Rasps consisted of highly variable broadband trains of 2 to 336 ticks that lasted between 74 and 10 500ms. The 260
rasps that were measured contained frequencies between 344 and 34 000 Hz with an average peak frequency of
3409 Hz. The frequency structure of ticks within rasps was highly variable and included both positive and negative
trends. This finding makes sablefish one of the few deep-sea fish for which sounds have been validated and
described. The documentation of sablefish sounds will enable the use of passive acoustic monitoring methods in fish-
eries and ecological studies of this commercially important deep-sea fish. V
C2020 Acoustical Society of America.
https://doi.org/10.1121/10.0001071
(Received 5 December 2019; revised 17 March 2020; accepted 24 March 2020; published online 16 April 2020)
[Editor: Arthur N. Popper] Pages: 2295–2301
I. INTRODUCTION
Fish sounds have been studied since at least the late
1800 s (Dufoss
e, 1874) and since then there have been numer-
ous accounts of the variability that exists in fish sound pro-
duction (Fish, 1948;Fish et al., 1952;Moulton, 1963;
Schneider, 1966;Tavolga, 1971;Hawkins, 1993;Kaatz,
2002;Ladich, 2004;Fine and Parmentier, 2015;Zeyl et al.,
2016). The ability to recognize fish sounds is becoming
increasingly useful for passive acoustic monitoring (PAM)
studies on population and ecosystem health (Rountree et al.,
2006;Slabbekoorn et al.,2010;Riera et al., 2016;Archer
et al., 2018;Lindseth and Lobel, 2018). In order to use PAM,
examples of sounds from each species of fish need to be vali-
dated and available for comparison to the sounds recorded
through PAM (Rountree et al.,2002). There are currently
34 300 known fish species (Froese and Pauly, 2019)and
sound production has been reported for fewer than 1000 spe-
cies (Lobel et al.,2010), although an updated number remains
to be confirmed. This number is growing as new fish sounds
are being described (Wilson et al., 2004;Riera et al., 2018;
Rountree et al.,2018). Despite these efforts, the capacity for
sound production remains to be investigated for the majority
of fish species (Rountree et al.,2002,2019).
There is increasing interest in understanding the dynam-
ics and health of deep-sea ecosystems such as sponge reefs
(Archer et al., 2018), seamounts (Department of Fisheries
and Oceans Canada, 2011), and banks, as these systems are
fragile and vulnerable to overfishing (Koslow et al., 2000).
The soundscape of the deep sea is poorly known and the use
of PAM methods to study these ecosystems is becoming
more common (Rountree et al., 2012;Wall et al., 2014).
The deep-sea sablefish (Anoplopoma fimbria, order
Scorpaeniformes, family Anoplopomatidae), also known as
black cod, is an economically important ground fish native
to the North Pacific Ocean ranging from Baja California to
the Bering Sea and throughout the Aleutian Islands into
waters off the Kamchatka Peninsula, Russia, and northern
Japan (Wilkins and Saunders, 1997;Jacobson et al., 2001).
Adult sablefish inhabit the upper continental slope and deep
continental shelf at depths of 200–1280 m (Wilkins and
Saunders, 1997;Jacobson et al., 2001). Sablefish support
valuable commercial and recreational fisheries in Alaska
(Warpinski et al., 2016), Japan, Russia, and along the U.S.
West Coast (Koslow et al., 2000). In addition, thanks to its
high growth rate and market value, sablefish aquaculture is
developing in several countries, including the U.S. and
Canada (Sumaila et al., 2007;Sanchez-Serrano et al., 2014;
National Marine Fisheries Service, 2017). The sablefish was
first suggested to produce sounds in an unpublished study of
captive fish by Meldrim (1965) and later based on deep-sea
recordings associated with sablefish presence at deep-sea
observatories (Sirovic et al., 2012), but these observations
have not been substantiated. Confirmation of sablefish sound
production, together with a validated description of sablefish
sound characteristics, would provide researchers with a new
tool to monitor the species using passive acoustics.
The goal of this study was to determine if sablefish pro-
duce sounds, and if so, to provide validated sound descrip-
tions to enable future PAM studies of the species. Captive
sablefish were observed and recorded both in an open-water
aquaculture facility and in a sablefish research station.
II. MATERIALS AND METHODS
A. Data collection
Acoustic recordings were obtained at two facilities:
Golden Eagle Sablefish Farm (GESF) (BC, Canada), where
a)
This paper is part of a Special Issue on The Effects of Noise on Aquatic
Life.
b)
Electronic mail: ariera@uvic.ca
J. Acoust. Soc. Am. 147 (4), April 2020 V
C2020 Acoustical Society of America 22950001-4966/2020/147(4)/2295/7/$30.00
ARTICLE
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a few hundred sablefish were held in 30 m
2
net pens, and the
NOAA Northwest Fisheries Science Center at Manchester
Research Station (NWFSC-MRS) (WA, USA), where 20–30
sablefish were held in 3.66 m diameter tanks. There were
only mature adult sablefish at GESF [size range: 46–85 cm
total length (TL)]. At the NWFSC-MRS adult sablefish (size
range: 35–75 cm TL) were monitored from seven tanks, and
juveniles (size range: 3–5 cm TL) from a single tank. The
adult sablefish at NWFSC-MRS were distributed in four
tanks with mixed sexes, one tank with only males, and two
tanks with unknown sexes.
At both facilities, sablefish were monitored for sound
production in real time. Recordings were made at 96 kHz
(24 bit), to a Zoom-H1 recorder (Zoom North America,
Hauppauge, NY) with an uncalibrated SQ26-01 hydrophone
(sensitivity ¼193.5 dB re: 1 V/lPa, Cetacean Research
Technology, Seattle, WA). At the NWFSC-MRS, water
pumps were turned off in order to reduce noise.
A Song Meter SM4 recorder (Wildlife Acoustics,
Maynard, MA) with an HTI hydrophone (sensitivity
¼165 dB re: 1 V/lPa, High Tech Inc., Long Beach, MS)
was also deployed in a tank containing juvenile sablefish at
NWFSC-MRS to collect data on a continuous duty cycle at
96 kHz (16 bit) for up to 4 days. No alterations were made to
the regular schedules of pumps and filters for SM4 recordings.
B. Data post-processing
Acoustic measurements of selected parameters of all
sablefish sounds were made in Raven Pro 1.5 acoustic soft-
ware (Center for Conservation Bioacoustics, 2014) follow-
ing Charif et al. (2010). Recordings were visually inspected
in their entirety to identify sablefish sounds. Spectrograms
were displayed 15 s at a time with frequencies between 0
and 11 kHz [2825 fast Fourier transform (FFT), Hann win-
dow, 85% overlap]. Selection boxes were drawn around
each sound to measure the sound duration, the lowest peak,
and highest frequency, the 5th and 95th percentile frequen-
cies (F. 5% and F. 95%, respectively), and bandwidth 90%
(BW 90%) (Charif et al., 2010). Raven Pro automatically
computed these values based on the selection boundaries. F.
5% is the frequency that divides the selection into two fre-
quency intervals containing 5% of the energy at the bottom
and 95% of the energy at the top, while F. 95% is the fre-
quency separating 95% of the energy at the bottom and 5%
at the top. BW 90% is the difference between F. 5% and F.
95% frequencies. The peak frequency is the frequency at
which maximum power occurs within the signal. For each
variable, the measurements reported include minimum,
maximum, and mean 6SE (standard error).
Sablefish sounds are comprised of a number of broad-
band ticks that are separated from each other by variable
durations. To differentiate between one sound and the next,
an arbitrary cutoff of 1 s was used.
A subset of 72 sablefish sounds from the NWFSC-MRS
was used to count the number of ticks per sound and mea-
sure tick-specific duration and frequency parameters (724
FFT, Hann window, 85% overlap). The duration between
ticks, or period, was calculated as the time between the
beginning of one tick and the beginning of the next tick
(Fig. 1). The inter-tick interval was calculated as the time
between the end of one tick and the beginning of the next
tick. The tick repetition rate was calculated by dividing the
number of ticks in a given sound by the duration of that
sound. Within-sound variation in tick frequency structure
(F. 5%, peak, F. 95%, and BW 90%) was tested for correla-
tion with elapsed time for 57 unique sounds having 8 or
more ticks. Spearman Rank correlation on log transformed
frequencies was performed due to non-linear data trends
using SAS/STAT software (SAS Institute Inc., 2012).
Mm. 1. Audio clip of short sablefish rasp (with fewer than
eight ticks) corresponding to the spectrogram displayed
in Fig. 1. This is a file of type “WAV” (561 KB).
III. RESULTS
Sounds attributed to adult sablefish were produced by
highly agitated fish that displayed aggressive behavior
(charging and nipping the hydrophone) during net pen trans-
fer at GESF. Similar sounds were recorded from captive
specimens at the NWFSC-MRS but were not associated
with any specific behavior.
Sounds were recorded at GESF between 2:00 pm and
6:00 pm. Sounds were recorded at the NWFSC-MRS in 5 C
water between 7:00 am and 4:30 pm. At NWFSC-MRS, two
or more rasps were heard in each of the four tanks that con-
tained mixed genders, and the tank that had only males. No
rasps were positively identified in the recordings from the
two tanks with unknown genders nor in the tank that con-
tained juveniles.
The duration of sablefish sounds ranged between 74 and
10 493 ms (average of 1342 696 SE; Table I) and they
FIG. 1. Waveform (top) and spectrogram (bottom) of a short rasp (with
fewer than eight ticks) produced by sablefish (Anoplopoma fimbria) at the
Northwest Fisheries Science Center in Manchester (1800 FFT Hann win-
dow with 85% overlap). The temporal measurements are illustrated: rasp
duration (a), tick duration (b), and period (c). A clip of the sound is avail-
able as a multimedia file (Mm. 1).
2296 J. Acoust. Soc. Am. 147 (4), April 2020 Riera et al.
https://doi.org/10.1121/10.0001071

consisted of highly variable trains of 3 to 336 ticks (average
30 65, Table II).
Due to the similarity of these sounds with cetacean
rasps (Marrero P
erez et al., 2017), they were subsequently
referred to as “rasps.” Rasps were highly variable in dura-
tion, number of ticks, and frequency structure (Figs. 1and
2). Rasp frequency ranged from 344 to 33 968 Hz, with an
average peak frequency of 3409 Hz 6118 (Table I).
Additional frequency- and time-based measurements of
sablefish rasps are presented in Table I.
Mm. 2. Audio clip of long sablefish rasp (with more than
eight ticks) corresponding to the spectrogram displayed
in Fig. 2(A). This is a file of type “WAV” (843 KB).
Mm. 3. Audio clip of long sablefish rasp (with more than
eight ticks) corresponding to the spectrogram displayed
in Fig. 2(B). This is a file of type “WAV” (840 KB).
Mm. 4. Audio clip of long sablefish rasp (with more than
eight ticks) corresponding to the spectrogram displayed
in Fig. 2(C). This is a file of type “WAV” (1543 KB).
In addition to inter-rasp frequency variation, the inspec-
tion of individual ticks uncovered wide intra-rasp frequency
variation (Table II). Some rasps were made of ticks whose
bandwidth remained relatively constant throughout the entire
call (e.g., the tick with the greatest bandwidth was only about
400 Hz higher than the tick with the smallest bandwidth).
Other rasps presented bandwidth variability among their ticks
as great as 27.5 kHz. For some rasps, the bandwidth was
greater for the first few ticks, and then became narrower as the
call progressed [e.g., Figs. 1and 2(A)]. Most rasps exhibited
significant positive correlations between one or more tick fre-
quency measures and elapsed time within the rasp (see supple-
mentary Table I in the supplemental material
1
). Examples of
both significant positive and negative trends in tick frequency
withinaraspareshowninFig.3.
The duration of ticks ranged between 1 and 53 ms, with
an average of 11 ms 60.1 (Table II). The period ranged
between 0.2 and 64.3 ms, with an average of 6 ms 60.1
(Table II). Within the same rasp, the period varied as little
as 0.2 ms (in a rasp with 3 ticks) and as much as 62.7 ms
(in a rasp with 23 ticks).
IV. DISCUSSION
The analysis of the recordings collected at both loca-
tions revealed a total of 260 broadband high-frequency
sounds (average 3 KHz peak) referred to as rasps. These
sounds were composed of a series of short (average 11 ms),
broadband tick sounds that varied in frequency content and
time-interval between successive ticks (period). These char-
acteristics match the description of the sounds reported by
Meldrim (1965) from his unpublished study on captive
sablefish, and also support the hypothesis that sablefish
could have been the source of the broadband pulses recorded
by Sirovic et al. (2012) in Barkley Canyon. The attribution
of the rasp sounds to sablefish was supported by independent
observations in two different facilities. Real-time
TABLE I. Acoustic variables for sablefish (Anoplopoma fimbria) rasps recorded at two facilities, showing stats for each of them and both pooled together.
SE ¼standard error of the mean. Min ¼minimum. Max ¼maximum. F. ¼Frequency. BW ¼Bandwidth. F. 5% is the frequency that divides the signal into
two frequency intervals containing 5% and 95% of the energy in the signal. F. 95% is the frequency that divides the signal into two frequency intervals con-
taining 95% and 5% of the energy in the signal. BW 90% is the difference between the 5% and 95% frequencies.
Acoustic variables
GESF (N¼152) Manchester Research Station (N¼108) Pooled (N¼260)
Min Max Mean (6SE) Min Max Mean (6SE) Min Max Mean (6SE)
Low F. (Hz) 535 11 668 2446 6111 344 4817 1826 699 344 11 668 2188 679
F. 5% (Hz) 551 11 766 2552 6113 773 5859 23586109 551 11 766 2471 680
Peak F. (Hz) 574 12 258 3086 6144 434 9234 38636192 434 12 258 3409 6118
F. 95% (Hz) 1816 13 090 5418 6182 2203 30 305 94936581 1816 30 305 7111 6291
High F. (Hz) 2053 13154 6549 6209 2395 33 968 11 362 6716 2053 33 968 8548 6353
BW 90% (Hz) 375 8988 2866 6150 891 25 711 71366528 375 25 711 4640 6269
Duration (ms) 74 4323 732 652 98 10 493 22016192 74 10 493 1342 696
TABLE II. Acoustic variables for the sablefish (Anoplopoma fimbria) ticks
recorded at the Manchester Research Station (N¼2136 except for the
period and inter-tick interval where N¼2064, and the tick repetition rate
where N¼72). Ticks are the broadband pulses that make up the rasps.
These ticks were measured from a sub-sample of 72 rasps. SE ¼standard
error of the mean. Min ¼minimum. Max ¼maximum. F. ¼Frequency.
BW ¼Bandwidth. F. 5% is the frequency that divides the signal into two
frequency intervals containing 5% and 95% of the energy in the signal. F.
95% is the frequency that divides the signal into two frequency intervals
containing 95% and 5% of the energy in the signal. BW 90% is the differ-
ence between the 5% and 95% frequencies.
Acoustic variables Min Max Mean (6SE)
Low F. (Hz) 401 22 140 2570 639
F. 5% (Hz) 797 22 406 3178 639
Peak F. (Hz) 1066 23801 5398 662
F. 95% (Hz) 2133 32180 10 540 6102
High F. (Hz) 2481 41 463 12 225 6114
BW 90% (Hz) 363 28 852 7362 696
Duration (ms) 1 53 11 60.1
Number of ticks/rasp 3 336 30 65
Period (ms) 0.2 64.3 6 60.1
Inter-tick interval (ms) 0 63 5 60.1
Tick repetition rate (number of ticks/s) 5 63 18 61
J. Acoust. Soc. Am. 147 (4), April 2020 Riera et al. 2297
https://doi.org/10.1121/10.0001071

observations at GESF indicated that an artificial source of
the sounds was highly unlikely, though the possibility of
other biological sources could not be ruled out in the open
water pens. However, recordings of the same type of sounds
in tanks of adult sablefish at the NWFSC-MRS facility
confirmed sablefish as the only possible source. The fact
that rasps were recorded in multiple tanks with adults but
were absent from other tanks further reduces the likelihood
that they were artifacts.
This newly validated description of sablefish sounds
suggests that PAM surveys for sablefish can be used both in
fisheries applications and in studies of deep-sea ecology in
areas within the species’ geographic range.
Sablefish is one of the top 10 key commercial species in
the U.S., with an important fishery in the North Pacific Region
(Alaska) and Pacific Region (California, Oregon,
Washington), where the total annual landings revenue was
between 102 and 185 10
6
U.S. dollars between 2006 and
2015 (National Marine Fisheries Service, 2017). In British
Columbia, there have been concerns about the sablefish stock
declining below a sustainable yield, and management strate-
gies have been designed to promote stock growth while
FIG. 2. Three examples of sablefish rasps illustrating the high variation in rasp
structures and variation in tick frequency structure produced by sablefish
(Anoplopoma fimbria) at the Northwest Fisheries Science Center in Manchester.
Each example includes waveform (top) and spectrogram (bottom) (1800 FFT
Hann window with 85% overlap). A clip of each sound is available as multime-
dia files (Mm. 2–Mm. 4). (A) Rasp with a trend for increasing F.5. (Mm. 2).
The top panel is an expansion of the first tick in the middle panel, delineated
with a box. (B) Rasp with relatively constant tick frequency structure (Mm. 3).
(C) Long rasp with high variation in tick frequency structure (Mm. 4).
FIG. 3. (Color online) Examples of two rasps exhibiting significant correla-
tions of tick frequency parameters (5% frequency: square, peak frequency:
triangle, 95% frequency: circle) with the elapsed time from the beginning
of the rasp. The Spearman Rank Correlation (r) is indicated for 95% fre-
quency (top), peak frequency (middle), and 5% frequency (bottom) with
asterisks representing its significance level (* ¼0.05, ** ¼0.01,
*** ¼0.001, ns ¼not significant). (Top) Positive correlation (rasp ID 39 in
the supplementary table in the supplemental material
1
). (Bottom) Negative
correlation (rasp ID 48 in the supplementary table in the supplemental
table
1
).
2298 J. Acoust. Soc. Am. 147 (4), April 2020 Riera et al.
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attempting to maintain the economic performance (Cox et al.,
2011). Stock biomass is currently assessed via trawling sur-
veys and fishery catch data (Wilkins and Saunders, 1997;
Koslow et al., 2000;Warpinski et al., 2016). The use of PAM
has the potential to enhance current sablefish management by
providing another independent monitoring tool.
The sablefish fishery in the Gulf of Alaska suffers great
reductions in catches due to sperm whale (Physeter macro-
cephalus) and killer whale (Orcinus orca) depredation on
longline fishing gear (Peterson and Hanselman, 2017;Wild
et al., 2017). An acoustic decoy has been used to broadcast
vessel-hauling noises known to attract whales at a distance
away from the vessel performing true hauls, thus reducing
the number of interactions between whales and fishing ves-
sels (Wild et al., 2017). It would be interesting to investigate
the response of whales to sablefish sounds. If whales are
attracted to rasps, perhaps adding recordings of sablefish
rasps to the vessel-hauling sounds could increase the effi-
cacy of the decoy as an attractant.
The soundscape of the deep-sea is poorly known, and
fish sounds have been described for very few deep-sea spe-
cies (see reviews in Rountree et al., 2012;Wall et al., 2014;
Parmentier et al., 2018). This limited knowledge could be
due to a series of factors including the need for specialized
equipment, inaccessibility, the non-continuous nature of fish
sound production (they might not be vocal at the moment of
recording), and the low amplitude of fish sounds that makes
them susceptible to masking and reduces their detection
range (Rountree et al., 2012;Wall et al., 2014). The results
presented here add sablefish as one of the few demonstrated
cases of sound production in deep-sea fishes. Knowing what
sablefish sound like will also facilitate a more complete
understanding of events that are already being monitored
with video at underwater cabled observatories (Doya et al.,
2014) where concurrent acoustic recordings are available.
This study demonstrates that sablefish produce sounds, and
therefore this knowledge is useful for PAM studies. How and
why the fish make the sound (if there is a specific function) is
unknown, and what follows is a discussion of some options.
The mechanism by which sablefish produce sounds is
currently unknown. The phylogenetic relationships of sable-
fish to other scorpaeniform fishes is uncertain, but the family
Anoplopomatidae is currently thought to be most closely
related to the greenlings (Hexagrammidae) and sculpins
(Cottidae) (Imamura and Yabe, 2002;Shinohara and
Imamura, 2007;Nelson et al., 2016). Unfortunately, despite
the high diversity of sculpins, sounds have only been
described in two genera (see reviews in Zeyl et al., 2016;
Bolgan et al., 2019) and it is unknown in greenlings.
The broadband high-frequency rasps produced by sable-
fish are highly unusual among fish, and previously unknown
for any scorpaeniform fish (Bolgan et al., 2019). High fre-
quency fish sounds have been reported for Clupeiformes
(Wilson et al., 2004;Rountree et al., 2018), Cypriniformes
and Salmoniformes (Rountree et al., 2018), Perciformes such
as grunts (Bertucci et al.,2014) and cichlids (Lanzing, 1974;
Nelissen, 1978;Kottege et al., 2015;Spinks et al., 2017),
Siluriformes (Ghahramani et al., 2014;Mohajer et al., 2015),
and Gadiformes (Vester et al., 2004). An important distinction
between the high frequency sounds produced by sablefish and
those produced by other fishes, is that in most other known
cases, the sound production mechanism involves the gas blad-
der (Tavolga, 1971;Ladich, 2004) which is absent in sablefish
(Nelson et al., 2016). In fish that lack a swim bladder, the most
common sound-producing mechanism is stridulation, which
consists of rubbing hard body parts together, such as bones,
teeth, or fin spines (Tavolga, 1971;Ladich, 2004). The high
variation in sablefish rasp frequency is consistent with a stridu-
latory mechanism (Fine and Parmentier, 2015). For the sculpin
species whose sound production has been described, average
peak frequency was between 50 and 500 Hz (Zeyl et al., 2016),
which is much lower than that of sablefish ticks (5398 662 Hz;
Table II). The tick duration for cottid fishes was also shorter
than that of sablefish; an average of 30 64ms to 68612 ms
(Zeyl et al., 2016)comparedto1160.1 ms (Table II).
High frequency stridulatory sounds can also be found in
some catfish (Ghahramani et al., 2014;Mohajer et al.,2015),
grunt (Bertucci et al., 2014), and cichlid (Lanzing, 1974;
Nelissen, 1978;Kottege et al.,2015;Spinks et al.,2017)spe-
cies. The average peak frequency for catfish has been reported
to be between 521 6240 Hz and 2895 6276 Hz (Parmentier
et al., 2010), while the average peak frequency for grunts was
718 6180 Hz (Bertucci et al.,2014). Sounds produced by
grunts also consisted of a series of units that were themselves
composed of a variable number of pulses (Bertucci et al.,2014).
In sablefish, frequency parameters vary greatly between ticks
within the same rasp (Fig. 3), but how the frequency of each
unit varies within the series is not described for grunts, making
comparisons difficult. One of the biggest differences between
sablefish rasps and the cichlid high-frequency sounds is the
number of components; cichlids have calls composed of an
average of two pulses (Spinks et al.,2017), whereas sablefish
raspshaveanaverageof30andupto300ticksperrasp.This
difference translates into an overall longer duration for rasps.
Another less well-known sound production mechanism
found in some scorpaeniform species uses a novel
“chordophone” mechanism involving vibrations of tendons
to achieve higher frequencies than possible through muscle
contraction alone (see review in Bolgan et al., 2019). Future
research is needed to determine if sablefish sounds are pro-
duced by a stridulatory, chordophone, or other mechanism.
Although the lack of a swim bladder precludes an air
movement sound production mechanism (see review in
Rountree et al., 2018) in sablefish, a superficial similarity to
Pacific herring (Clupea pallasii) “fast repetitive tick” (FRT)
sounds has implications for PAM applications. Pacific herring
sounds are also composed of long trains of up to 65 ticks
(Wilson et al., 2004). The durations of rasps and FRTs are also
comparable, ranging between 0.7 and 10.5 s (average 1.3 s) for
rasps and 0.6 and 7.6 s (average 2.6 s) for FRTs (Wilson et al.,
2004). However, the period for rasps was highly variable (pre-
senting no clear pattern), whereas the period for herring FRTs
usually increases or decreases at a steady rate (Wahlberg and
Westerberg, 2003;Wilson et al., 2004;Kuznetsov, 2009).
J. Acoust. Soc. Am. 147 (4), April 2020 Riera et al. 2299
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Most fishes where hearing has been examined hear best
around 200 Hz (Mann et al., 2007) and have audibility thresh-
olds up to 3 kHz (Ladich and Fay, 2013) but sablefish rasps
can get up to 30 kHz and whether they can hear their own
sounds remains unknown. The ability to produce sounds is not
necessarily associated with a matching sensitivity to hear
them (Ladich, 2000), so an inability to hear the rasps does not
preclude the possibility of other functions such as predator
avoidance. However, high frequency hearing exists for some
fishes in the subfamily Alosinae, which have been reported to
hear ultrasounds from 40 to 80 kHz (Mann et al.,2001).
Those Alosinae species can also hear the lower frequency
components of sounds, down to 200 Hz (Mann et al.,2001),
which indicates that the ability to hear ultrasounds does not
rule out the ability to hear low frequencies. All fishes can
detect particle motion through the otolith organs, but their
ability to perceive sound pressure could be limited to the pres-
ence of gas-filled structures (Hawkins and Popper, 2018),
which are absent in sablefish (Nelson et al.,2016). Sablefish
rasps have a mean peak of 3409 6118 Hz (Table I), which
falls within the range of hearing thresholds of hearing special-
ists (Ladich, 2000), so it is possible they have evolved a simi-
lar hearing specialist ability through an unknown mechanism
not involving the gas bladder. The hearing abilities of sable-
fish need to be investigated, and if possible, such studies
should design methodologies that produce data that are com-
parable between species and laboratories (Popper et al.,2019).
The skilfish, Erilepis zonifer, is the only other species
in the family Anoplopomatidae (Froese and Pauly, 2019). A
few studies have been conducted on the distribution and
biology of the skilfish (Zolotov et al., 2014), but no data is
available regarding their possible sound production. The
capacity for sound production is often shared by species of
the same family (Wall et al., 2014;Spinks et al., 2017;
Parmentier et al., 2018), which makes the skilfish a good
candidate for further studies to verify the hypothesis.
Although sound production in sablefish has been demon-
strated, it remains unclear if the sounds are produced for an
acoustic function such as intra-species communication which
requires an unexpected ability to hear high frequency sounds,
an inter-species signal that aids in predator avoidance which
does not require hearing sensitivity, or is entirely incidental to
some unknown physiological function. Regardless, the
description of sablefish sounds provides scientists with the
opportunity to use PAM methodologies in the study and man-
agement of the species. In addition, even if entirely incidental,
determination of the physiological mechanism that produces
such unusual sounds would be informative in and of itself,
and suggests that PAM could be used to monitor spatial and
temporal patterns in that physiological process. Future work
could include studies on hearing, sound production mecha-
nism, and behaviours associated with vocal activity.
ACKNOWLEDGMENTS
The authors would like to thank Hannah Britton-Foster
and Kelsie Murchy for their assistance in the field. Thanks
are also due to the crew at the GESF (Terry Brooks,
Quinten, Dave, Trevor, Mike, Anthony) and personnel at the
Manchester Research Station (Rick Goetz and Cortney
Jensen). We are grateful to MERIDIAN for their support.
Funding was provided by DFO Contribution Agreements, an
NSERC Discovery grant, CFI and BCKDF equipment
grants, and the Liber Ero Foundation.
1
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